http://www.collembola.org/taxa/tullidae.htm - Last updated on 2024.09.24 by Frans Janssens
Checklist of the Collembola: Tullbergiidae

Familia Tullbergiidae Bagnall RS, 1935:238 key  key  key  key

Genus Number
of species
Ameritulla3
Anaphorura1
Boudinotia1
Clavaphorura1
Delamarephorura5
Dinaphorura16
Doutnacia4
Fissuraphorura7
Jevania2
Karlstejnia5
Marcuzziella1
Mesaphorura53
Metaphorura11
Mexicaphorura1
Mixturatulla1
Multivesicula7
Najtiaphorura4
Neonaphorura14
Neotullbergia5
Paratullbergia11
Pongeiella5
Prabhergia3
Psammophorura3
Rotundiphorura1
Scaphaphorura5
Sensilatullbergia1
Stenaphorura8
Stenaphorura21
Tasphorura1
Tillieria2
Tullbergia43
Tullbergiella2
Wankeliella7
Weinera1
Tullbergiidae236
Genera with number of species
  Syn.: Collembola Onychiuridae Kovacevic, M, Amorim, MJB, Hackenberger, BK et Scott-Fordsmand, JJ, 2022:3
  Syn.: Onychiuridae Onychiurinae Kovacevic, M, Amorim, MJB, Hackenberger, BK et Scott-Fordsmand, JJ, 2022:5


  Syn.: Onychiuridae Tullberginae(sic) Salmon, JT, 1943:377 ISS
  Syn.: Onychiuridae Tullberginae(sic) Mari Mutt, JA et Bellinger, PF, 1990:55 ISS
  Syn.: Onychiuridae Tullberginae(sic) Palacios-Vargas, JG et Díaz, M, 1996:166 ISS
  Syn.: Onychiuridae Tullberginae(sic) Skidmore, RE, 1995:53 ISS
  Syn.: Onychiuridae Tullberginae(sic) Mari Mutt, JA et Bellinger, PF, 1996:169 ISS
  Syn.: Onychiuridae Tullberginae(sic) González, V, Díaz, M et Palacios-Vargas, JG, 1997:46 ISS

Tullbergiidae from the USA
Body narrow worm-shaped
2012.02.16 © Kouri, J.
Tullbergiidae from the UK
Pigmentation absent, eyes absent
2012.10.02 © Murray, A.

  Syn.: Collembola Tuubergiinae(sic) Kaprus, IJ et Goblyk, KM, 2015:8 ISS
  Syn.: Poduromorpha Tullbergidae(sic) Qazi, F et Shayanmehr, M, 2016:127 ISS
  Syn.: Poduromorpha Tullbergidae(sic) Özata, MA, Sevgili, H et Kaprus, IJ, 2017:247 ISS
  Syn.: Collembola Tullbergidae(sic) Berg, M, van Langevelde, F, Wynhoff, I et van Bezouw, R, 2018:29 ISS
  Syn.: Poduromorpha Tullbergidae(sic) Mandal, GP, 2018:116 ISS
  Syn.: Collembola Tullbergidae(sic) Mandal, GP, 2018:207 ISS
  Syn.: Collembola Tullbergiinae(sic) Ni, Z, Yan, X-M, Chang, L-A, Sun, X, Wu, D-H et Zhang, B, 2020:8 lapsus
Tullbergiidae from Holland
Antennae short, legs short
2020.09.28 © Kamsteeg, G.

  Syn.: Poduromorpha Tullbergidae(sic) Cucini, C, Fanciulli, PP, Frati, F, Convey, P, Nardi, F et Carapelli, A, 2021:12 ISS
  Syn.: Poduromorpha Tullbergidae(sic) Cipola, NG, Queiroz, GC et Betsch, J-M, 2022:944 ISS

Subfamilia Clavaphorurinae

Remark by Janssens, F. 2012.11.04: For genera with 4 sensory clubs in the ant.3 sense organ we tentatively propose the subfamily Clavaphorurinae with type genus Clavaphorura Salmon, 1943.




Genus Clavaphorura Salmon, JT, 1943:377

Species septemseta Salmon, JT, 1943:377, t.t.

Clavaphorura septemseta from New Zealand
After Salmon JT, 1943 Pl.41 Fig.22
            Syn.: Tullbergia septemspina Schaller, 1949 nn




Subfamilia Tullbergiinae Deharveng L, 2004:427

  Syn.: Tullbergiinae Group I Bagnall, RS, 1935:240
  Syn.: Tullbergiinae Tullbergiini Bagnall, 1935



Genus Tullbergia Lubbock, 1876

  Syn.: ... Börneria(sic) Willem, 1902 IOS
  Syn.: Tullbergiidae Protullbergia Bagnall, 1947
  Syn.: Tullbergiinae Bachatullbergia Wray, 1958
  Syn.: Tullberginae Chaetophorura Rusek, 1976
  Syn.: Tullbergiinae Granuliphorura Rusek, 1976


Tullbergia sp. from Tasmania
Post-antennal organ and pseudocellus
After Ireson JE, 1990 Fig.70
Tullbergia sp. from Tasmania
Ant.3 organ with 3 sensory clubs
After Ireson JE, 1990 Fig.71

  Syn.: Onychiuridae (Tullbergiinae) Tullberia(sic) Greenslade, P et Rusek, J, 1996:33 ISS
Tullbergia sp. from Antarctica
2003 © Stevens, M.
Remark by ICZN. 2007:841: Placed on the Official List of Names in Zoology. Opinion 291.

Tullbergia? sp. from Tasmania
2014.04.04 © Murray, A.
Tullbergia? sp. from Tasmania
Anal spines
2014.04.04 © Murray, A.
Tullbergia sp. from Antarctica
After Fox D, 2020
2020 © Siwanowicz, I.





Species africana Martynova, 1979

Species alcirae Palacios-Vargas, JG et Salazar-Martínez, AE, 2014:24

Species anops Christiansen, K et Bellinger, P, 1980:482

Species antarctica Lubbock, 1876, t.t.

Tullbergia antarctica from Antarctica
After Lubbock J, 1879 Pl.XI Fig.1

Species arctica Wahlgren, 1900

Tullbergia arctica from the USA
After Fjellberg A, 1984 Fig.6B
Tullbergia arctica from the USA
2023.10.28 © Wang, G.
Tullbergia arctica from the USA
2023.10.28 © Wang, G.
Tullbergia arctica from the USA
2023.10.28 © Wang, G.
Tullbergia arctica from the USA
2023.10.28 © Wang, G.
Tullbergia arctica from the USA
Ant.3 sense organ with 3 external sensilla
2023.10.28 © Wang, G.
Tullbergia arctica from the USA
Abd.4-6
Anal spines
2023.10.28 © Wang, G.
Tullbergia arctica from the USA
Abd.4-6
Anal spines
2023.10.28 © Wang, G.
Tullbergia arctica from the USA
Pseudocellus striate
2023.10.28 © Wang, G.
Tullbergia arctica from the USA
Th.1 with 1+1 pseudocelli
2023.10.28 © Wang, G.
Tullbergia arctica from the USA
Th.2-3 with 2+2 pseudocelli
2023.10.28 © Wang, G.
Tullbergia arctica from the USA
Abd.1-4 with 2+2 pseudocelli
Abd.5 with 1+1 pseudocelli
2023.10.28 © Wang, G.
Tullbergia arctica from the USA
Ant.4 sensilla as thin as acuminate setae
2023.10.28 © Wang, G.

Species australica Womersley, 1933

Species bella Fjellberg, A, 1988:37

Tullbergia bella from Norway
After Fjellberg, A, 1988 Fig.2G
            Syn.: Tullbergia sp. Fjellberg, 1984



Species bisetosa Börner, 1903

            Syn.: Tullbergia insularis Womersley, H, 1935:212
            Syn.: Tullbergia insularis Salmon, JT, 1964:146
            Syn.: Tullbergia insularis Mari Mutt, JA et Bellinger, PF, 1990:55
            Syn.: Mesaphorura sp. Watson, 1967


Species collis Bacon, GA, 1914:84

Tullbergia collis from the USA
After Bacon GA, 1914 Fig.1
Tullbergia collis from the USA
After Bacon GA, 1914 Fig.2

Species crozetensis Deharveng, 1981

Species duops Christiansen, K et Bellinger, P, 1980:489

             Syn.: Tullbergia doups(sic) Cutz-Pool, LQ et Vázquez-González, M, 2012:109,ca ISS

Species gambiensis Womersley, H, 1935:211

            Syn.: Tullbergia gambiense(sic) Womersley, H, 1935:211 IOS
            Syn.: Tullbergia scalpellata Salmon, JT, 1949:17
            Syn.: Tullbergia subantarctica Salmon, JT, 1949:18
            Syn.: Tullbergia tillyardi Womersley, H, 1939:317
            Syn.: Tullbergia trisetosa Womersley, H, 1939:130


             Syn.: Tullbergia gambiense(sic) Womersley, 1939 ISS
             Syn.: Tullbergia gambiense(sic) Salmon, JT, 1964:146 ISS
             Syn.: Tullbergia gambiense(sic) Lawrence, 1968 ISS
             Syn.: Tullbergia gambiense(sic) Greenslade, P in Houston, WWK, 1994:62 ISS
             Syn.: Tullbergia gambiense(sic) Greenslade, P et Rusek, J, 1996:33 ISS
             Syn.: Tullbergia gambiense(sic) Najt, J et Weiner, WM in Najt, J et Matile, L, 1997:12 ISS



Species ghilarovi (Khanislamova, 1987) Dunger, W & Schlitt, B, 2011:144

Species harti (Rusek, 1991) Waltz & Hart, 1997

Species huetheri da Gama, 1968

            Syn.: Tullbergia hütheri(sic) da Gama, 1968 IOS

Species inconspicua de Izarra, DC, 1965:265

Species insularis Wahlgren, E, 1906:8

Species kilimanjarica (Delamare Deboutteville, C, 1953:818) Paclt, 1959

Species latens Christiansen, K et Bellinger, P, 1980:497

Species mala Christiansen, K et Bellinger, P, 1980:500

Tullbergia mala from the USA
After Fjellberg A, 1984 Fig.6A
Remark by Simón Benito, J.C. 2006.10.16: This species should be revised, it does not belong in Mesaphorura (pseudocelli are not star shaped), also not in Chaetophorura. It is a species complex, with 2-3 sensory rods in the third antennal segment.
Tullbergia mala from the USA
2023.12.23 © Wang, G.
Tullbergia mala from the USA
2023.12.23 © Wang, G.
Tullbergia mala from the USA
2023.12.23 © Wang, G.
Tullbergia mala from the USA
Pseudocellus striate
2023.12.23 © Wang, G.
Tullbergia mala from the USA
Ant.3 organ with 2 distinctly large curved sensilla
2023.12.23 © Wang, G.

Species massoudi

            Syn.: Tullbergia massoudi Hermosilla & Rubio, 1976 nn

Species maxima Deharveng, 1981

Species mediantarctica Wise, 1967

Tullbergia mediantarctica from Antarctica
Dorsal aspect
After Fox D, 2020
2020 © Siwanowicz, I.
Tullbergia mediantarctica from Antarctica
Ventral aspect
After Fox D, 2020
2020 © Siwanowicz, I.
Tullbergia mediantarctica from Antarctica
Lateral aspect
After Fox D, 2020
2020 © Siwanowicz, I.

Species meridionalis Cassagnau & Rapoport, 1962

            Syn.: Tullbergia meridionalis f. bisetosa Cassagnau & Rapoport, 1962


Tullbergia meridionalis from Chile
After Rapoport EH & Rubio IR, 1963 Fig.73
Tullbergia meridionalis from Argentina
After Arbea JI, 2016 Fig.20-21

Species meridionalis2 Barra, J-A, 1995:132

Species mexicana Handschin, 1928

Species minensis Arlé, R, 1960:165

Species mixta Wahlgren, E, 1906:8

            Syn.: Tullbergia mediantarctica Yue & Tamura, 2001
            Syn.: Tullbergia sp. Bulavintsev, 1990 =?
            Syn.: Tillieria penai Weiner & Najt, 1994


Species nearctica Bernard, EC, 2016:472

Tullbergia nearctica from the USA
After Bernard EC, 2016 Fig.13AB

Species nulla Christiansen, K et Bellinger, P, 1980:502

Species obtusochaeta (Rusek, 1976) Christiansen, K & Bellinger, P, 1980:504

             Syn.: Tullbergia obtusichaeta(sic) Bernard, EC, 2016:477 ISS

Species ouatilou Najt, J et Weiner, WM in Najt, J et Matile, L, 1997:11

Species paranensis Izarra, 1969

Species pomorskii (Smolis, 2010)

Species quadrisetosa (Willem, 1902) Börner, 1902

Börneria(sic) quadrisetosa from Antarctica
After Willem V, 1902 Pl.I Fig.1
             Syn.: Tullbergia quadriseta(sic) Womersley, H, 1935:212 ISS


Species rapoporti Arbea, JI, 2016:65

Tullbergia rapoporti from Argentina
After Arbea JI, 2016 Fig.1-2
Tullbergia rapoporti from Argentina
After Arbea JI, 2016 Fig.18

Species schaefferi Salmon, 1974

Species templei Wise, KAJ, 1970:210

            Syn.: Tullbergia mediantarctica Lugg, Johnstone & Griffin, 1978 cf
            Syn.: Tullbergia mixta Watson, 1967


Species tolanara Thibaud, J-M, 2008:510

Species trisetosa Schäffer, 1897

            Syn.: Aphorura trisetosa ssp. quadrisetosa Salmon, JT, 1964:146
             Syn.: Aphorura trisetosa ssp. quadriseta(sic) Womersley, H, 1935:212 ISS
            Syn.: Tullbergia tullbergi Womersley, 1939


Species vancouverica (Rusek, 1976) Christiansen, K & Bellinger, P, 1980:508

             Syn.: Tullbergia vancourverica(sic) Poole, RW in Poole, RW et Gentili, P, 1997:54 ISS

Species ventanensis Rapoport, 1963

Species womersleyi (Bagnall, 1947) Greenslade P in Houston, 1994:63


Genus Anaphorura Izarra, 1972

Species lavadoi (Izarra, 1972) Palacios-Vargas & Díaz, 1995:171, t.t.

             Syn.: Anaphorura lavadori(sic) Palacios-Vargas & Díaz, 1995 ISS



Genus Fissuraphorura Rusek, 1991

Species sp.1 Barra, J-A, 1998:526

Species cubanica Rusek, 1991:145

             Syn.: Fissuraphorura Cubanica(sic) Palacios-Vargas & Díaz, 1996 ISS

Species deharvengi Rusek, 1991

Species duplex (da Gama, 1962) Lucianez & Simon, 1992

Species gisini (Selga, 1963) Lucianez & Simon, 1992

Species miscellanea Barra, J-A, 1995:135

Species selgae (Prabhoo, 1971) Lucianez & Simon, 1992


Genus Tillieria Weiner, WM et Najt, J in Chazeau, J et Tillier, S, 1991:123 key

  Syn.: ... Tilleria(sic) Greenslade P, [1996] ISS

Species araucariensis Weiner, WM et Najt, J in Chazeau, J et Tillier, S, 1991:124

Species insularis Weiner, WM et Najt, J in Chazeau, J et Tillier, S, 1991:124, t.t.


Genus Dinaphorura Bagnall, RS, 1935:241

  Syn.: Onychiuroidea (Tullbergiidae) Austraphorura Bagnall, 1947
  Syn.: Onychiuridae (Tullbergiinae) Astraphorura(sic) Rodgers, D et Greenslade, P, 1996:1370 ISS


Species americana Rapoport, 1962:17




            Syn.: Tullbergia spinosissima Cassagnau & Rapoport, 1962


Species chazeaui Weiner, WM et Najt, J in Chazeau, J et Tillier, S, 1991:126

Species diversispina Womersley, H, 1935:213

            Syn.: Dinaphorura harrisoni Bagnall, 1947
            Syn.: Tullbergia septemspina Schaller, 1949 nn =?
            Syn.: Protullbergia willemi Bagnall, 1947


Species harrisoni Bagnall, 1947

Species jarai Najt & Rubio, 1978

Species kerguelensis Deharveng, 1981


Dinaphorura laterospina from New Zealand
After Salmon, J.T. 1941 Pl.48 Fig.132.
Species laterospina Salmon, JT, 1941:311

Species magellanica Rubio, 1974

Species matileorum Najt, J in Tillier, S, 1988:29

Species najtae Arbea, JI, 2016:78

Dinaphorura najtae from Argentina
After Arbea JI, 2016 Fig.36-37

Species nerudai Arbea, JI, 2016:75

Dinaphorura nerudai from Argentina
After Arbea JI, 2016 Fig.27-28

Species novaezealandiae Womersley, H, 1935:213

            Syn.: Dinaphorura novae-zealandeae(sic) Womersley, H, 1935:213 IOS
             Syn.: Dinaphorura novae-zealandiae(sic) Salmon, JT, 1941:310 ISS
             Syn.: Dinaphorura novae-hollandiae(sic) Womersley ISS
             Syn.: Dinaphorura novae-hollandae(sic) Womersley ISS
             Syn.: Dinaphorura novae-zealandeae(sic) Salmon, JT, 1964:147 ISS
             Syn.: Dinaphorura nova-zealandiae(sic) Najt, J in Tillier, S, 1988:32 ISS
             Syn.: Dinaphorura novaezealandeae(sic) Rodgers, D et Greenslade, P, 1996:1369 ISS



Species pefauri Rubio & Najt, 1979

Species spinosissima (Wahlgren, E, 1906:9) Bagnall, RS, 1935:242, t.t.

            Syn.: Tullbergia septemspina Schaller, 1949 nn =?
            Syn.: Dinaphorura diversispina Greenslade, P, 2015:9 laps


Species tooheyensis Rodgers, D et Greenslade, P, 1996:1374

Species wahlgreni (Bagnall, 1947) Najt & Rubio, 1978, s.d.


Genus Neonaphorura Bagnall, RS, 1935:240 key  key

  Syn.: Onychiuridae Tullbergia Denis, J-R, 1932:366
  Syn.: Tullbergia Bachatullbergia Wray, 1958
  Syn.: Tullbergiinae Bachytullbergia(sic) Salmon, JT, 1964:149 ISS
  Syn.: Tullbergiidae Spinaphorura Arbea, 1987 =i



Neonaphorura sp. from France
2006 © Boldsystems.org
Neonaphorura sp. from France
2006 © Boldsystems.org

Species adulta (Gisin, 1944) Salmon, JT, 1964:149

            Syn.: Tullbergia (Neonaphorura) duboscqi f. adulta Gisin, 1944
             Syn.: Neonaphorura adula(sic) Traser, G et Thibaud, J-M, 1999:106 ISS

Species alicatai Giuga & Jordana, 2013

Species ariegica Arbea, JI, 1991:179

Species duboscqi (Denis, J-R, 1932:366) Bagnall, RS, 1935:241, t.t.

            Syn.: Tullbergia Duboscqi(sic) Denis, J-R, 1932:366 IOS
Tullbergia Duboscqi(sic) from France
After Denis J-R, 1932 Fig.IV.1


            Syn.: Neonaphorura anglicana Bagnall, 1936
             Syn.: Neonaphorura anglicanus(sic) Salmon, JT, 1964:149 ISS
            Syn.: Tullbergia duboscqi f. adulta Gisin, 1944



             Syn.: Neonaphorura duboscqui(sic) Bagnall, RS, 1935:241,tt ISS
             Syn.: Tullbergia dubosqi(sic) Haarlon, 1957 ISS
             Syn.: Neonaphorura duboscqui(sic) Salmon, JT, 1964:149 ISS
             Syn.: Neonaphorura dubosqui(sic) Kopeszki, H et Meyer, E, 1996:225 ISS
             Syn.: Neonaphorura dubosqi(sic) ssp. dubosqi(sic) Traser, G et Thibaud, J-M, 1999:106 ISS
Neonaphorura duboscqi from the UK
Two large and 4 small anal spines
2013.04.27 © Murray, A.





Species dungeri Schulz, 1994

Species graffi Kaprus, IJ et Goblyk, KM, 2015:2

Neonaphorura graffi from Ukraine
After Kaprus IJ & Goblyk KM, 2015 Fig.1

Species hexaspina Arbea, JI et Mateos, E, 1991:169

Species howdeni (Wray, 1958) Salmon, JT, 1964:149

Species loksai Traser, G et Thibaud, J-M, 1999:105

Remark by Kaprus, I.J. & Goblyk K.M. 2015:1-2: Among Neonaphorura species cited by Dunger and Schlitt (2011), only N. loksai Traser et Thibaud, 1999 does not fully fit with the diagnosis of the subfamily Tullbergiinae, because antennal III sense organ has only two sensory clubs [therefore matching subfamily Mesaphorurinae]. However, other characters of N. loksai are typical for the genus Neonaphorura.
Species moravica (Rusek, 1966) Khanislamova, 1988

Species navarrensis Arbea, JI, 1991:184

Species novemspina (Gisin, 1963) Christian, 1987

            Syn.: Tullbergia septemspina Schaller, 1949 nn =?


Neonaphorura novemspina from Holland
Abd.6 with 8 (paired) spines in focus (*)
2019.12.25 © Berg, M.
Det.: Berg, M.
Neonaphorura novemspina from Holland
Abd.6 with 9th (unpaired) spine in focus (*)
2019.12.25 © Berg, M.
Det.: Berg, M.

Species ortali Palissa, 2006

Species zakarpatica Kaprus, IJ et Goblyk, KM, 2015:4

Neonaphorura zakarpatica from Ukraine
After Kaprus IJ & Goblyk KM, 2015 Fig.12


Genus Tasphorura Greenslade, P et Rusek, J, 1996:28

Species vesiculata Greenslade, P et Rusek, J, 1996:28, t.t.


Genus Stenaphorura Absolon, K, 1900:411

  Syn.: Tullbergiidae Tullbergia Börner, 1902, nec Lubbock, 1876
  Syn.: Aphorurini(sic) Mesaphorura Börner, C, 1901:3 =?
  Syn.: Tullbergiinae Mesaphorura Bagnall, RS, 1935:236
  Syn.: Tullbergiini Stenaphorurella Luciañez & Simón, 1992


Species sp. Greenslade, P et Rusek, J, 1996:33

Species denisi Bagnall, 1935, B

            Syn.: Tullbergia parisi Denis, 1943
             Syn.: Stenaphorura parisii(sic) Stach, 1954 ISS
             Syn.: Stenaphorura parisii(sic) Salmon, JT, 1964:152 ISS


             Syn.: Stenaphorurella Denisi(sic) Khanislamova G M, Sterzynska M et Weiner W M, 1997:503 ISS
Stenaphorura denisi from the UK
2012.11.28 © Murray, A.
Stenaphorura denisi from the UK
2012.11.28 © Murray, A.
Stenaphorura denisi from the UK
Two large and two small anal spines; in front two bumps (*)
2012.11.28 © Murray, A.
Stenaphorura denisi from the UK
2012.11.28 © Murray, A.
Stenaphorura denisi from the UK
2013.01.11 © Murray, A.
Stenaphorura denisi from the UK
Two large and two small anal spines; in front two bumps
2013.01.11 © Murray, A.
Stenaphorura denisi from the UK
2013.01.11 © Murray, A.

             Syn.: Stenaphorura denisii(sic) Shaw, P, 2017:147 ISS
Stenaphorura denisii(sic) from the UK
After Shaw, P in Ware, C. & al, 2017 p.147 right



Species gibsoni Murphy, 1965

Species japygiformis Absolon, K, 1900:412, t.t.

             Syn.: Stenaphorura iapygiformis(sic) Börner, C, 1901:9 ISS
             Syn.: Stenaphorura iapygiformis(sic) Börner, C, 1901:24 ISS
             Syn.: Stenaphorura iapygiformis(sic) Bagnall, RS, 1935:241,tt ISS
             Syn.: Tullbergia iapygiformis(sic) Bagnall, RS, 1935:237 ISS
             Syn.: Stenaphorura iapygiformis(sic) Bagnall, 1935 ISS
             Syn.: Tullbergia japygidiformis(sic) Salmon, 1964 ISS

Species lubbocki Bagnall, 1935

Species metaparisi (Traser & Weiner, 1999) Dunger, W & Schlitt, B, 2011:138

Species quadrispina Börner, C, 1901:699, B

            Syn.: Aphorura 4-spina(sic) ssp. taurica Lebedinsky, 1904
            Syn.: Stenaphorura axelsoni Bagnall, 1935
            Syn.: Stenaphorura absoloni Bagnall, 1936


Stenaphorura quadrispina (bar=2µm)
Hexagonal epicuticular ultrastructure with secondary granules
After Nickerl J & al, 2012 Fig.2C'
Stenaphorura quadrispina (bar=500nm)
Epicuticular secondary granule
After Nickerl J & al, 2012 Fig.2C''

Species shaconage Bernard, EC, 2016:467

Stenaphorura shaconage from the USA
After Bernard EC, 2016 Fig.10AB


Subfamilia Mesaphorurinae Dunger, W & Schlitt B, 2011:19

  Syn.: Tullbergiinae Stenaphorurini Luciáñez & Simón, 1992
  Syn.: Tullbergiidae Stenaphorurinae Deharveng, L, 2004:427
Mesaphorurinae juv. from Canada
2014.02.02 © Hochhalter, R.
Mesaphorurinae from the USA
Pseudocellus at base of antenna
2020.06.12 © Speer, J.H.
Mesaphorurinae from the USA
Cuticular tubercles
2020.06.12 © Speer, J.H.
Mesaphorurinae from the USA
Ant.3 organ with 2 sensory clubs (*)
and 3 guard setae
2020.06.19 © Speer, J.H.
Mesaphorurinae from the USA
Post-antennal organ with 2 rows of 12 vesicles
2020.06.19 © Speer, J.H.





Genus Mesaphorura Börner, C, 1901:1 key  key

  Syn.: Tullbergiidae Tullbergia Börner, 1902, nec Lubbock, 1876
  Syn.: Collembola Folsomia(sic) Moore, JC, Saunders, P, Selby, G, Horton, H, Chelius, MK, Chapman, A et Horrocks, RD, 2005:117 lapsus


  Syn.: Aphorurini(sic) Mesaphorum(sic) Börner, C, 1901:24 ISS
  Syn.: Aphorurini(sic) Mesaphorum(sic) Börner, C, 1901:233 ISS
  Syn.: Collembola Mesophorura(sic) Kopeszki, H et Meyer, E, 1996:233 ISS
Mesaphorura sp. juv. from the UK
2013.01.12 © Murray, A.
Mesaphorura sp. juv. from the UK
2013.01.12 © Murray, A.
Mesaphorura sp. from Belgium
2017.07.16 © Huskens, M.-L.
Mesaphorura sp. from the USA
Abd.6 with crescentic ridges and anal spines on strong papillae
2020.02.28 © Speer, J.H.
Mesaphorura sp. from Canada
2021.11.26 © Brousseau, P.-M.
Mesaphorura sp. from Canada
With 2 anterior + 2 posterior cephalic pseudocelli
2021.11.26 © Brousseau, P.-M.
Mesaphorura sp. from Canada
Abd.6 with crescentic ridges and anal spines
2021.11.26 © Brousseau, P.-M.
Mesaphorura sp. from Canada
Anterior pseudocellus (pso); post-antennal organ (PAO) with 2 parallel rows of vesicles
2021.11.26 © Brousseau, P.-M.

  Syn.: Onychiuridae(sic) Mesaprophorura(sic) Kovacevic, M, Amorim, MJB, Hackenberger, BK et Scott-Fordsmand, JJ, 2022:4 ISS


Species sp.1 Janssens F sensu Thibaud, 1993

            Syn.: Mesaphorura sp.cf.italica Thibaud, 1993
            Syn.: Mesaphorura sp.cf.italica Mari Mutt, JA et Bellinger, PF, 1996:169


Species sp.2 Janssens F sensu Palacios-Vargas & Díaz, 1996

            Syn.: Mesaphorura sp.ca.collis Palacios-Vargas, JG et Díaz, M, 1996:171


Species sp.3 Janssens F sensu Palacios-Vargas & Díaz, 1996

            Syn.: Mesaphorura sp.ca.pacifica Palacios-Vargas, JG et Díaz, M, 1996:172


Species amazonica Oliveira & Thibaud, 1992

Species arbeai Simón Ruiz Martinez & Lucianez, 1994

Species atlantica (Rusek, 1979) Thibaud, J-M & Palacios-Vargas, JG, 1999:29

Species baconae Bagnall, 1947

Species bassolsae Palacios-Vargas, JG et Díaz, M, 1996:12

Species betschi Rusek, 1979, B

            Syn.: Mesaphorura massoudi Rusek, 1979


             Syn.: Tullbergia betchi(sic) Fjellberg, A, 1988:39 ISS

Species choudhurii Yosii, 1966

Mesaphorura choudhurii from India
After Mandal GP & Hazra AK, 2009 Fig.15a+b

             Syn.: Mesaphorura choudhuri(sic) Mandal, GP et Hazra, AK, 2009:73 ISS
             Syn.: Mesaphorura choudhuri(sic) Mandal, GP et Hazra, AK, 2009:77 ISS
             Syn.: Mesaphorura choudhuri(sic) Mandal, GP et Hazra, AK, 2009:115 ISS
             Syn.: Mesaphorura choudhuri(sic) Mandal, GP et Hazra, AK, 2009:126 ISS
             Syn.: Mesaphorura choudhuri(sic) Mandal, GP et Hazra, AK, 2009:128 ISS


Species critica Ellis, WN, 1976:230

            Syn.: Mesaphorura krausbaueri Ardanaz, 1986, partim


Mesaphorura critica from Australia
After Greenslade P, 1992 Fig.2
Mesaphorura critica from China
After Gao Y & Bu Y, 2022 Fig.32

Species delamarei Weiner, 1991

Species dubia Börner, 1902

Species florae Simón, Ruiz, Martinez & Lucianez, 1994

             Syn.: Mesaphorura phlorae(sic) García-Gómez, A, Castaño-Meneses, G et Palacios-Vargas, JG, 2009:911 ISS

Species foveata Bonet, 1944

Species guyana Thibaud, 2004

Species himalayensis Yosii, 1971

Remark by Simón Benito, J.C. 2006.10.16:in litt.: This species should be revised, it does not belong in Mesaphorura (pseudocelli are not star shaped), but in Chaetophorura or Weinera.

Species hygrophila (Rusek, J, 1971:199) Ellis, 1976:232

             Syn.: Mesaphorura hygrophyla(sic) Ellis, 1976 ISS

Species hylophila Rusek, J, 1982:14, B

            Syn.: Mesaphorura krausbaueri Jordana & Beruete, 1983, partim


Mesaphorura hylophila from China
After Bu Y & Gao Y, 2017 Fig.1

Species intermedia Prabhoo, 1971

Species iowensis (Mills, HB, 1932:264) Bagnall, 1937

Species isochaeta Arbea, JI et Jordana, R, 1989:8

  Subspecies leitzaensis Jordana, 1993

Species italica (Rusek, J, 1971:203) Ellis, 1974:109, B

            Syn.: Mesaphorura krausbaueri Jordana & Beruete, 1983, partim
            Syn.: Mesaphorura krausbaueri Jordana & al., 1987, partim


             Syn.: Mesaphorura italic(sic) Kahrarian, M, 2019:42 ISS

Species jarmilae Rusek, J, 1982:20

Species javieri (Palacios-Vargas, JG & Díaz, M, 1996:18) Palacios-Vargas, JG in Sánchez-Ramos, G, Reyes-Castillo, P & Dirzo, R, 2005:364

Species jeremiei Thibaud, J-M et Weiner, WM in Najt, J et Matile, L, 1997:74

Mesaphorura jeremiei from New Caledonia
After Thibaud J-M & Weiner WM, 1997 Fig.34

Species jevanica Rusek, J, 1996:208, B

            Syn.: Mesaphorura sp. Rusek, 1996


Species jirii Rusek, J, 1982:18

Species krausbaueri Börner, C, 1901:2, t.t., B

            Syn.: Mesaphorura Krausbaueri(sic) Börner, C, 1901:2,tt IOS
            Syn.: Tullbergia arctica Linnaniemi, 1912 =?
            Syn.: Tullbergia Krausbaueri(sic) f. atypica II Denis, J-R, 1924:225
            Syn.: Tullbergia iowensis Mills, 1932
            Syn.: Paratullbergia macdougalli Bagnall, 1936
            Syn.: Mesaphorura baconae Bagnall, 1947
            Syn.: Mesaphorura krausbaueri var. bonariensis Rapoport, 1959


             Syn.: Mesaphorura Krausbaueri(sic) Absolon, K, 1901:389 ISS
             Syn.: Mesaphorura Krausbaueri(sic) Absolon, K, 1901:580 ISS
             Syn.: Tullbergia Krausbaueri(sic) Denis, J-R, 1924:226 ISS
             Syn.: Tullbergia Krausbaueri(sic) Denis, J-R, 1924:227 ISS
             Syn.: Tullbergia Krausbaueri(sic) Loksa, I, 1961:224 ISS
             Syn.: Tullbergia krausboueri(sic) Salmon, JT, 1964:150 ISS
             Syn.: Tullbergia kraubaueri(sic) Wibo, C, 1971:107 ISS
             Syn.: Mesaphorura krausberi(sic) Hopkin, 1997 ISS
             Syn.: Folsomia(sic) krausberi(sic) Moore, JC, Saunders, P, Selby, G, Horton, H, Chelius, MK, Chapman, A et Horrocks, RD, 2005:117 ISS
             Syn.: Mesaphorura krausbauerii(sic) Thibaud, J-M, 2008:510,gr ISS

  Subspecies inermis Axelson, WM, 1905:790


            Syn.: Tullbergia arctica Kloet, GS et Hincks, WD, 1945:5 =?




Species macrochaeta Rusek, 1976, B

            Syn.: Metaphorura affinis f. B Ellis, WN, 1976:225
            Syn.: Mesaphorura krausbaueri Jordana & Beruete, 1983, partim
            Syn.: Mesaphorura krausbaueri Arbea & Jordana, 1985, partim
            Syn.: Mesaphorura krausbaueri Jordana & al., 1987, partim
            Syn.: Tullbergia krausbaueri Petersen, H, 1971:133
            Syn.: Tullbergia krausbaueri Greenslade, 1992 gr
            Syn.: Mesaphorura (krausbaueri-group) sp. Greenslade, 1990


Mesaphorura macrochaeta from Australia
After Greenslade P, 1992 Fig.1
             Syn.: Mesaphorura macrocheta(sic) D'Haese, CA, 2003:570 ISS
             Syn.: Mesaphorura macrocheta(sic) D'Haese, CA, 2003:397 ISS
             Syn.: Mesaphorura macrocheata(sic) Moore, JC, Saunders, P, Selby, G, Horton, H, Chelius, MK, Chapman, A et Horrocks, RD, 2005:117 ISS
             Syn.: Mesaphorura macrocheata(sic) Rochefort, S, Therrien, F, Shetlar, DJ et Brodeur, J, 2006:63 ISS
Mesaphorura macrochaeta (bar=2µm)
Hexagonal epicuticular ultrastructure with secondary granules
After Nickerl J & al, 2012 Fig.5D

Mesaphorura macrochaeta from the UK
2012.11.15 © Murray, A.
Mesaphorura macrochaeta from the UK
2012.11.15 © Murray, A.
Mesaphorura macrochaeta from the UK
2012.11.15 © Murray, A.
Mesaphorura macrochaeta from the UK
2012.11.15 © Murray, A.
Mesaphorura macrochaeta from Holland
Prominent anal spines and macrosetae on abd.6
2013.11.27 © van Bezouw, R.

            Syn.: Mesaprophorura(sic) macrochaeta Kovacevic, M, Amorim, MJB, Hackenberger, BK et Scott-Fordsmand, JJ, 2022:4
Mesaphorura macrochaeta from the USA
2023.03.29 © Wang, G.
Mesaphorura macrochaeta from the USA
With atypical dark colouration
2023.07.30 © Wang, G.
Mesaphorura macrochaeta from the USA
With atypical dark colouration
2023.07.30 © Wang, G.
Mesaphorura macrochaeta from the USA
Post-antennal organ
2023.07.30 © Wang, G.



Species mani Thibaud, 2002

Species maricaensis Fernandes, LH et de Mendonça, MC, 2004:17

Species matilei Thibaud & Palacios-Vargas, 2000

Species minutissima Salmon, JT, 1944:141

Species occidafricana Murphy, 1965

Species orousseti Najt, Thibaud & Weiner, 1990

            Syn.: Mesaphorura longispina Arbea, JI et Jordana, R, 1991:118
            Syn.: Mesaphorura longispina ssp. stellae Arbea, JI et Selga, D in Jordana, R et Ariño, AH, 1994:76


Species pacifica Rusek, 1976

Mesaphorura pacifica from China
After Bu Y & Gao Y, 2017 Fig.1O

Species petterdassi (Fjellberg, A, 1988:38) Sterzynska & Ehrnsberger, 1998

Tullbergia petterdassi from Norway
After Fjellberg, A, 1988 Fig.3A

Species pongei Rusek, 1982, B

Species redondoi Jordana, R et Arbea, JI in Jordana, R et Ariño, AH, 1994:4

Mesaphorura redondoi from Spain
After Jordana R & Arbea JI, 1994 Fig.2a
Remark by Dányi, L. & Traser, G. 2008:28: Jordana et al. (1997) considered this taxon as a subspecies of Mesaphorura isochaeta Arbea & Jordana, 1990, endemic of the Iberian Peninsula. However, this status is questionable in the light of the Hungarian occurences.


Species rudolfi Rusek, 1986

Species ruseki (Christiansen & Bellinger, 1980) Miranda Rangel & Palacios-Vargas, 1992


Species schembrii Thibaud, J-M et Christian, E, 1989:76

Species sensibilis Rusek, J, 1973:79

Species simoni Jordana, R et Arbea, JI in Jordana, R et Ariño, AH, 1994:2

Mesaphorura simoni from Spain
After Jordana R & Arbea JI, 1994 Fig.1a

Species simplex (Gisin, 1958)

Species spelaea (Nosek & Neuherz, 1976)

Species subitalica Thibaud, 1996

Species sylvatica (Rusek, J, 1971:195) Dunger, W, 1977:180

             Syn.: Mesaphorura sylavtica(sic) Kopeszki, H et Meyer, E, 1996:234 ISS
             Syn.: Mesaphorura silvatica(sic) Kaprus, IJ, Pomorski, RJ, Skarzynski, D et Potapov, MB, 2005:893 ISS
Mesaphorura sylvatica from Russia
Lateral aspect; size = 0.4mm
After Panina IV & al, 2019 Fig.1A
Mesaphorura sylvatica from Russia
Dorsal aspect
After Panina IV & al, 2019 Fig.1C
Mesaphorura sylvatica ♀ from Russia
Ventral aspect
After Panina IV & al, 2019 Fig.1B
Mesaphorura sylvatica ♀ from Russia
Internal morphology
Lateral internal aspect
After Panina IV & al, 2019 Fig.2A
Mesaphorura sylvatica ♀ from Russia
Internal morphology
Digestive and excretory system
After Panina IV & al, 2019 Fig.4AB
Mesaphorura sylvatica ♀ from Russia
Internal morphology
Nervous system
After Panina IV & al, 2019 Fig.4CD
Mesaphorura sylvatica ♀ from Russia
Internal morphology
Reproductive system
After Panina IV & al, 2019 Fig.4EF
Mesaphorura sylvatica ♀ from Russia
Internal morphology
Abdominal muscles lateral internal view
After Panina IV & al, 2019 Fig.6A
Mesaphorura sylvatica ♀ from Russia
Internal morphology
Remnant furcal extensor muscle (red)
Modified after Panina IV & al, 2019 Fig.6A



Species tenuisensillata Rusek, 1974, B

             Syn.: Mesaphorura tenuisensillana(sic) Kaprus, IJ, Pomorski, RJ, Skarzynski, D et Potapov, MB, 2005:897 ISS

Species thalassophila Bagnall, 1937

Species tuberosa Bonet, 1944

Species yosiii (Rusek, J, 1967:191) Rusek, 1973, B

            Syn.: Mesaphorura krausbaueri Arbea & Jordana, 1985, partim
            Syn.: Mesaphorura krausbaueri Jordana & al., 1987, partim


             Syn.: Tullbergia (Mesaphorura) yosii(sic) Rusek, J, 1971:192 ISS
             Syn.: Mesaphorura yosii(sic) Rusek, 1973 ISS
             Syn.: Mesaphorura yosii(sic) Rusek, 1979 ISS
             Syn.: Mesaphorura yosii(sic) Ponge in Pesson, 1980 ISS
             Syn.: Mesaphorura yossii(sic) Palacios-Vargas et al., 1982 ISS
             Syn.: Mesaphorura yosii(sic) Hågvar, 1983 ISS
             Syn.: Mesaphorura yosii(sic) Ponge, J-F, 1983:363 ISS
             Syn.: Mesaphorura yosii(sic) Hågvar, 1984 ISS
             Syn.: Mesaphorura yosii(sic) Hågvar & Abrahamsen, 1984 ISS
             Syn.: Mesaphorura yosii(sic) Pozo, 1986 ISS
             Syn.: Tullbergia yosii(sic) Takeda, 1987 ISS
             Syn.: Mesaphorura yosii(sic) Kuznetsova, NA, 1988:28 ISS
             Syn.: Tullbergia vosii(sic) Leinaas, HP, 1988:278 ISS
             Syn.: Mesaphorura yosii(sic) Luciañez Sanchez, MJ et Simon Benito, JC, 1988:115 ISS
             Syn.: Mesaphorura yosii(sic) Szeptycki & Weiner in Razowski, 1990 ISS
             Syn.: Mesaphorura yosii(sic) Arbea, JI et Jordana, R, 1991:116 ISS
             Syn.: Mesaphorura yosii(sic) Najt, J, Thibaud, J-M et Jacquemart, S (+), 1991:165 ISS
Mesaphorura yosiii from Australia
After Greenslade P, 1992 Fig.3

             Syn.: Mesaphorura yosii(sic) Ponge, J-F, 1993:242 ISS
             Syn.: Mesaphorura yosii(sic) Barra, 1994 ISS
             Syn.: Mesaphorura yosii(sic) Greenslade, P in Houston, WWK, 1994:59 ISS
             Syn.: Mesaphorura yosii(sic) Thibaud, J-M et Lee, B-H, 1994:41 ISS
             Syn.: Mesaphorura yosii(sic) Thibaud, J-M, Najt, J et Jaquemart(sic), S (+), 1994:200 ISS
             Syn.: Mesaphorura yosii(sic) Barra, 1995 ISS
             Syn.: Mesaphorura yosii(sic) Oliveira, EP et Deharveng, L, 1995:369 ISS
             Syn.: Mesaphorura yossi(sic) Sterzynska, M et Kuznetsova, N, 1995:148 ISS
             Syn.: Mesaphorura yosii(sic) Kopeszki, H et Meyer, E, 1996:225 ISS
             Syn.: Mesophorura(sic) yosii(sic) Kopeszki, H et Meyer, E, 1996:233 ISS
             Syn.: Mesaphorura yosii(sic) Rusek, J, 1996:214 ISS
             Syn.: Mesaphorura yosii(sic) Hopkin, 1997 ISS
             Syn.: Mesaphorura yosii(sic) Najt, J et Weiner, WM in Najt, J et Matile, L, 1997:10 ISS
             Syn.: Mesaphorura yosii(sic) Thibaud, J-M et Weiner, WM in Najt, J et Matile, L, 1997:76 ISS
             Syn.: Tullbergia (Mesaphorura) yosii(sic) Liu, Y-Q, Hou, D-B et Li, Z-C, 1998:127 ISS
             Syn.: Mesaphorura yosii(sic) Ponge, J-F, 1999:58 ISS
             Syn.: Mesaphorura yosii(sic) Palacios-Vargas, JG, 2000:29 ISS
             Syn.: Mesaphorura yosii(sic) Ponge, J-F, 2000:510,B ISS
             Syn.: Mesaphorura yosii(sic) Park, K-H, Kim, K-G, Park, S-J, Kim, B-J et Lee, B-H, 2000:415 ISS
             Syn.: Mesaphorura yosii(sic) Sabatini & Innocenti, 2000 ISS
             Syn.: Mesaphorura yosii(sic) Peck, SB, 2001:103 ISS
             Syn.: Mesaphorura yosii(sic) Thibaud & Palacios-Vargas, 2001 ISS
             Syn.: Mesaphorura yosii(sic) Castaño-Meneses, G, Palacios-Vargas, JG et Cutz-Pool, L, 2004:138 ISS
             Syn.: Mesaphorura yosii(sic) Fernandes, LH et de Mendonça, MC, 2004:16 ISS
             Syn.: Mesaphorura yosii(sic) Gillet, S et Ponge, J-F, 2004:223,B ISS
             Syn.: Mesaphorura yosii(sic) Tamura & al., 2004 ISS
             Syn.: Mesaphorura yosii(sic) Janssens, F et Dethier, M, 2005:154,B ISS
             Syn.: Mesaphorura yosii(sic) Boitaud, L, Salmon, S Bourlette, C et Ponge J-F, 2006:451 ISS
             Syn.: Mesaphorura yosii(sic) Anichkin, AE, Belyaeva, NV, Dovgobrod, IG, Shveenkova, YB et Tiunov, AV, 2007:501 ISS
             Syn.: Mesaphorura yosii(sic) Fiera, C, 2007:18 ISS
             Syn.: Mesaphorura yosii(sic) Dányi, L et Traser, G, 2008:29 ISS
             Syn.: Mesaphorura yosii(sic) Thibaud, J-M, 2008:510 ISS
             Syn.: Mesaphorura yosii(sic) Xiong, Y, Gao, Y, Yin, W-Y et Luan, Y-X, 2008:730 ISS
             Syn.: Mesaphorura yosii(sic) Fernandes, LH, Nessimian, JL et de Mendonça, MC, 2009:1036 ISS
             Syn.: Mesaphorura yosii(sic) Nakamori, T, Fujiwara, N, Matsumoto, N et Okada, H, 2009:7 ISS
             Syn.: Mesaphorura yosii(sic) Busmachiu, G, 2010:153 ISS
             Syn.: Tullbergia yosii(sic) Suetsugu, N, Satoh, F et Koike, T, 2010:60 ISS
             Syn.: Mesaphorura yosii(sic) Chauvat, M, Trap, J, Perez, G, Delporte, P et Aubert, M, 2011:411 ISS
             Syn.: Mesaphorura yosii(sic) Dunger, W et Schlitt, B, 2011:86 ISS
             Syn.: Mesaphorura yosii(sic) Palacios-Vargas, JG, Castaño-Meneses, G et Estrada, DA, 2011:120 ISS
             Syn.: Mesaphorura yosii(sic) Peck, SB et Jacquemart, J, 2011:5 ISS
             Syn.: Mesaphorura yossii(sic) Cutz-Pool, LQ et Vázquez-González, M, 2012:109 ISS
             Syn.: Mesaphorura yosii(sic) Busmachiu, G, Bedos, A et Deharveng, L, 2015:101 ISS
             Syn.: Mesaphorura yosii(sic) Janion-Scheepers, C, Deharveng, L, Bedos, A et Chown, L, 2015:64 ISS
             Syn.: Mesaphorura yosii(sic) Thibaud, J-M, 2015:356 ISS
             Syn.: Mesaphorura yosii(sic) Thibaud, J-M, 2015:358 ISS
             Syn.: Mesaphorura yosii(sic) Thibaud, J-M, 2015:359 ISS
             Syn.: Mesaphorura yosii(sic) Bernard, EC, 2016:451 ISS
             Syn.: Mesaphorura yosii(sic) Bernard, EC, 2016:457 ISS
             Syn.: Mesaphorura yosii(sic) Bernard, EC, 2016:460 ISS
             Syn.: Mesaphorura yosii(sic) Bernard, EC, 2016:476 ISS
             Syn.: Mesaphorura yosii(sic) Yu, D-Y, Zhang, F, Stevens, MI, Yan, Q-B, Liu, M-Q et Hu, F, 2016:288 ISS
             Syn.: Mesaphorura yosii(sic) Bu, Y et Gao, Y, 2017:85 ISS
             Syn.: Mesaphorura yosii(sic) Bu, Y et Gao, Y, 2017:91 ISS
             Syn.: Mesaphorura yosii(sic) Bu, Y et Gao, Y, 2017:170 ISS
             Syn.: Mesaphorura yosii(sic) Palacios-Vargas, JG, Cortés-Guzmán, D et Alcocer, J, 2018:2 ISS
             Syn.: Mesaphorura yosii(sic) Babenko, A, Stebaeva, S et Turnbull, MS, 2019:34 ISS
             Syn.: Mesaphorura yosii(sic) Bu, Y et Gao, Y, 2019:85 ISS
             Syn.: Mesaphorura yosii(sic) Jiang, L-L, Dong, J, Jin, J-F et Zhang, F, 2019:104 ISS
             Syn.: Mesaphorura yosii(sic) da Silveira, TC et de Mendonça, MC, 2020:214 ISS
             Syn.: Mesaphorura yosii(sic) da Silveira, TC et de Mendonça, MC, 2020:215 ISS
             Syn.: Mesaphorura yosii(sic) Cucini, C, Fanciulli, PP, Frati, F, Convey, P, Nardi, F et Carapelli, A, 2021:9 ISS
             Syn.: Mesaphorura yosii(sic) Cucini, C, Fanciulli, PP, Frati, F, Convey, P, Nardi, F et Carapelli, A, 2021:12 ISS
             Syn.: Mesaphorura yosii(sic) Timmermans, MJTN, Arbea, JI, Campbell, G, King, MC, Prins, A et Kett, S, 2021:4 ISS
             Syn.: Mesaphorura yosii(sic) Cipola, NG, Queiroz, GC et Betsch, J-M, 2022:941 ISS
             Syn.: Mesaphorura yosii(sic) Gao, Y et Bu, Y, 2022:531 ISS
             Syn.: Mesaphorura yosii(sic) Gao, Y et Bu, Y, 2022:541 ISS
             Syn.: Mesaphorura yosii(sic) Gao, Y et Bu, Y, 2022:543 ISS
             Syn.: Mesaphorura yosii(sic) Godeiro, NN, Palacios-Vargas, JG, Gao, Y et Bu, Y, 2022:755 ISS
             Syn.: Mesaphorura yosii(sic) Li, J-Y, Godeiro, NN, Gao, Y, Bu, Y et Li, K, 2022:? ISS
             Syn.: Mesaphorura yosii(sic) Marcin, M, Raschmanová, N, Miklisová, D, Supinskė, J, Kanuk, J et Kovác, L, 2022:12 ISS
             Syn.: Mesaphorura yosii(sic) Marcin, M, Raschmanová, N, Miklisová, D, Supinskė, J, Kanuk, J et Kovác, L, 2022:20 ISS
             Syn.: Mesaphorura yosii(sic) Nguyen, ATT, Thuy, KN et Arbea, JI, 2022:47 ISS
             Syn.: Mesaphorura yosii(sic) Shveenkova, YB, 2022:5 ISS
             Syn.: Mesaphorura yosii(sic) Wang, B-F, Yin, J-Q, Wu, F-C, Wang, D-M, Jiang, Z-L et Song, X-Y, 2022:5 ISS
             Syn.: Mesaphorura yosii(sic) Jucevica, E, Grina, V et Melecis, V, 2023:258 ISS
             Syn.: Mesaphorura yosii(sic) Mehrafrooz Mayvan, M, Greenslade, P et Sadeghi-Namaghi, H, 2023:22 ISS
             Syn.: Mesaphorura yosii(sic) Mehrafrooz Mayvan, M, Greenslade, P et Sadeghi-Namaghi, H, 2023:84 ISS
Mesaphorura yosiii from the USA
Dorsal aspect
2023.07.27 © Wang, G.
Mesaphorura yosiii from the USA
Lateral aspect
2023.07.27 © Wang, G.
Mesaphorura yosiii from the USA
Ventral aspect
2023.07.27 © Wang, G.
Mesaphorura yosiii from the USA
Abd.6 with crescentic ridges
2023.07.27 © Wang, G.




Genus Neotullbergia Bagnall, RS, 1935:241

Remark by Dunger, W & Schlitt B. 2011:111: The genus Neotullbergia Bagnall needs a revision.
Remark by Janssens, F. 2012.11.04: The genus Neotullbergia Bagnall possibly is a genus complex given some species such as ramicuspis and crassicuspis have 2 sensory clubs while others such as staudacheri and tricuspis have 3 sensory clubs in the ant.3 sense organ.
Species americana Bonet, 1944

Species crassicuspis (Gisin, 1943) Salmon, JT, 1964:148, B

Species ramicuspis (Gisin, 1953) Salmon, 1964:148

            Syn.: Tullbergia tricuspis Strebel, 1957
            Syn.: Tullbergia tricuspis Paclt, 1961


Species staudacheri (Kos, 1940) Dunger, W & Schlitt, B, 2011:112

            Syn.: Tullbergia Neotullbergia tricuspis Gisin, 1944
            Syn.: Neotullbergia laingi Bagnall, 1936


Species tricuspis (Börner, 1902) Bagnall, RS, 1935:241, t.t.


Genus Metaphorura Stach, J, 1954:204 key

  Syn.: Tullbergiidae Tullbergia auctt. partim
  Syn.: Metaphorura Bagnall, 1936 u Metaphorura Bagnall, 1936 u Metaphorura Bagnall, 1936 u
  Syn.: Tullbergiidae Metaphorura Kloet, GS et Hincks, WD, 1945:5 u


Species affinis (Börner, 1902) Bagnall, 1936 sensu Stach, 1954, t.t.

            Syn.: Metaphorura börneri(sic) Bagnall, 1936 sensu Stach, 1954 IOS
             Syn.: Tullbergia bipartita ssp. borneri(sic) Bagnall, 1939 ISS
            Syn.: Metaphorura affinis type B Ellis, WN, 1976:227
            Syn.: Metaphorura bipartita? Skarzynski, D, Pomorski, RJ, Smolis, A, Weiner, WM, Szeptycki, A, Slawska, M et Sterzynska, M, 2002:32


Metaphorura affinis
2006 © Boldsystems.org
Metaphorura affinis from the UK
Two anal spines with medial conical projection
2013.04.27 © Murray, A.



Species bipartita (Handschin, 1920) Bagnall, 1936, t.t.

            Syn.: Metaphorura börneri(sic) Bagnall, 1936 IOS
             Syn.: Metaphorura börneri(sic) Bagnall, 1937 ISS
             Syn.: Metaphorura börneri(sic) Kloet, GS et Hincks, WD, 1945:5 ISS
             Syn.: Metaphorura börneri(sic) Denis, 1948 ISS
            Syn.: Tullbergia bagnalli Denis, 1943
             Syn.: Tullbergia Bagnalli(sic) Delamare Deboutteville, C et Jacquemart, S, 1961:8 ISS
            Syn.: Metaphorura affinis Stach, 1955




Species denisi Simón Benito, 1985

            Syn.: Metaphorura affinis f. C Ellis, WN, 1976:225


Species ellisi Simón Benito, 1985

            Syn.: Metaphorura affinis f. A Ellis, WN, 1976:225


Species incisa (Bonet, 1944) Castaño-Meneses, G, Palacios-Vargas, JG & Traser, G, 2000:1266

Species knowltoni (Wray, DL, 1950:97) Palacios-Vargas, JG & Díaz, M, 1996:173

Tullbergia knowltoni from the USA
After Wray DL, 1950 Pl.5 Fig.1

Species motuoensis Bu, Y et Gao, Y, 2017:87

Metaphorura motuoensis from Tibet
After Bu Y & Gao Y, 2017 Fig.1

Species orestia Pomorski, RJ, Skarzynski, D et Kaprus, I, 1998:265

Species riozoi Castaño-Meneses, G, Palacios-Vargas, JG et Traser, G, 2000:1263

Metaphorura riozoi from Iran
After Shayanmehr M & al, 2017 Fig.1a

Species triacantha (Börner, 1901) Dallai, R, Malatesta, E & Ramellini, P in Minelli, A, Ruffo, S & La Posta, S, 1995:8

             Syn.: Stenaphorura Ariacantha(sic) Börner, C, 1901:24 ISS
             Syn.: Stenaphorura 3-acantha(sic) Börner, C, 1901:24 ISS
             Syn.: Stenaphorura Ariacantha(sic) Börner, C, 1901:233 ISS

Species zhongi Bu, Y et Gao, Y, 2019:85

Metaphorura zhongi from Tibet
After Bu Y & Gao Y, 2019 Fig.1A



Genus Delamarephorura Weiner & Najt, 1999

  Syn.: Tullbergiidae Spicatella Thibaud, 2002


Species bedosae (Thibaud, 2002) Janion, Deharveng & Weiner, 2013

Species capensis Janion, Deharveng & Weiner, 2013

Species salti (Delamare Deboutteville, C, 1953:820) Weiner & Najt, 1999, t.t.

Species szeptyckii Barra & Weiner, 2009

Species tami Janion, Deharveng & Weiner, 2013


Genus Stenaphorura2 Thibaud, 2002 =i

Species marionae Thibaud, 2002


Genus Tullbergiella de Izarra, DC, 1965:267

Species allendei (Izarra, 1975) Mari Mutt & Bellinger, 1990:57

Species humilis (de Izarra, DC, 1965:267) Mari Mutt, JA & Bellinger, PF, 1990:57, t.t.

Tullbergia (Tullbergiella) humilis from Argentina
After de Izarra, 1965 Fig.12


Genus Karlstejnia Rusek, 1974

Species annae Rusek, 1974

Species montana Rusek, 1978

Species norvegica (Fjellberg, 1974) Potapov, M & Stebaeva, S, 1997:124

Species rusekiana Weiner, 1983

Species sibirica Rusek, 1978


Genus Prabhergia Salmon, 1965

Species imadatei Tamura H in Tamura, H et Zhao, L-J, 1996:791

Species indonesiae Yoshii, R et Suhardjono, YR, 1989:29

Species nayarii Salmon, 1965, t.t.


Genus Boudinotia Weiner, WM et Najt, J in Chazeau, J et Tillier, S, 1991:121

Species prima Weiner, WM et Najt, J in Chazeau, J et Tillier, S, 1991:121, t.t.


Genus Marcuzziella Rusek, 1975

  Syn.: Tullergiinae Marcuziella(sic) Petersen, H, 2011:486 ISS

Species tripartita Rusek, 1975


Genus Wankeliella Rusek, 1975 key

Species sp.1 Janssens F sensu Shveenkova YB, 2022

            Syn.: Wankeliella sp.n. aff. intermedia Shveenkova, YB, 2022:5


Species bescidica Smolis, A et Skarzynski, D, 2003:451

Species intermedia Potapov, M et Stebaeva, S, 1997:123

Species medialis Simón-Benito, JC et Jordana, R in Jordana, R et Ariño, AH, 1994:16

Wankeliella medialis from Spain
After Simón JC & Jordana R, 1994 Fig.1c

Species mediochaeta Rusek, 1975

Species peterseni Rusek, 1975 tt

Species pongei Rusek, 1978


Genus Paratullbergia Womersley, 1930 key

  Syn.: Tullbergiidae Tullbergia auctt. partim
  Syn.: Collembola Tullbergis(sic) Haybach, G, 1959:39 ISS
  Syn.: Collembola Tullbergis(sic) Haybach, G, 1959:Tab.VIII ISS


  Syn.: Tullbergiidae Paratullber gia(sic) Bu, Y et Gao, Y, 2019:64 ISS

Species brevispina Skarzynski & Pomorski, 1999

Species callipygos (Börner, 1902) Salmon, JT, 1964:147, B

            Syn.: Paratullbergia concolor Womersley, 1930 =?
            Syn.: Paratullbergia carpenteri Bagnall, 1935 =?
            Syn.: Paratullbergia womersleyi Bagnall, 1935
            Syn.: Tullbergia callipygos f. hirsuta Cassagnau, P, 1952:305
            Syn.: Tullbergiinae indet. Camacho & Prieto, 2011


             Syn.: Tullbergia callipygas(sic) Heungens, A et Van Daele, E, 1973:136,B ISS
             Syn.: Paratullbergia calliphygos(sic) Moore, JC, Saunders, P, Selby, G, Horton, H, Chelius, MK, Chapman, A et Horrocks, RD, 2005:117 ISS
Paratullbergia callipygos from the UK
2012.11.27 © Murray, A.
Paratullbergia callipygos from the UK
Elongate post-antennal organ
2012.11.27 © Murray, A.
Paratullbergia callipygos from the UK
Distinct large anal spines
2012.11.27 © Murray, A.

             Syn.: Tullbergia callipygosa(sic) Falahati HosseinAbad, A, Mehr, MS et Khyroodin, A, 2013:260 ISS
Paratullbergia callipygos from the UK
2013.04.05 © Murray, A.
Paratullbergia callipygos juv. from the UK
Transverse dorsal suture in 4th abdominal segment
2013.04.15 © Murray, A.
Paratullbergia callipygos from the UK
6th abdominal segment with ridges/bumps and large anal spines
2013.05.07 © Murray, A.
Paratullbergia callipygos juv. from Holland
Abd.4 with transverse suture
Abd.6 with ridges/bumps and 2 anal spines
2014.10.06 © van Duinen, J.
Paratullbergia callipygos juv. from the UK
Transverse dorsal suture in 4th abdominal segment
2014.12.28 © Nurcombe, E.
Paratullbergia callipygos from France
2019.10.04 © Brousseau, P.M.
Paratullbergia callipygos from Holland
Abd.6 with 2 anal spines
2022.11.12 © Kamsteeg, G.
Det.: Berg, M.
Paratullbergia callipygos from Holland
Abd.4 with transverse suture
2022.11.12 © Kamsteeg, G.
Det.: Berg, M.
Paratullbergia callipygos from Holland
Abd.6 with transverse ridges/bumps
2022.12.28 © Kamsteeg, G.
Det.: Berg, M.



Species caroli Lucianez Ortega & Simón, 1991

Species changfengensis Bu, Y et Gao, Y, 2015:56

Paratullbergia changfengensis from China
After Bu Y & Gao Y, 2015 Fig.1

Species chuana Gao, Y et Bu, Y, 2020:?

Species concolor Womersley, 1930, t.t.

Species indica Salmon, 1965

Species macdougalli Bagnall, 1936

Paratullbergia ?macdougalli? from France
2019.10.04 © Brousseau, P.M.
Paratullbergia ?macdougalli? from France
Abd.6 without large dorsal tubercles
2019.10.04 © Brousseau, P.M.
Paratullbergia ?macdougalli? from France
Abd.2+3: m3 absent
2019.10.04 © Brousseau, P.M.

Species qilianensis Bu, Y et Gao, Y, 2019:65

Species salmoni Prabhoo, 1971

             Syn.: Paratullbergia salmon(sic) Bu, Y et Gao, Y, 2015:60 ISS

Species trivandrana Prabhoo, 1971


Genus Multivesicula Rusek, 1982

Species aliciae Palacios-Vargas, JG et Díaz, M, 1996:15

Species ampla (Christiansen & Bellinger, 1980)

Tullbergia ampla from the USA
After Fjellberg A, 1984 Fig.5

Species columbica Rusek, 1982

             Syn.: Multivesicula columbia(sic) Gao, Y et Bu, Y, 2022:536 ISS

Species dolomitica Rusek, 1982

Species giljarovi Rusek, 1982

Species punctata Rusek, 1982

             Syn.: Multivesicula puctata(sic) Skidmore, RE, 1995:52 ISS

Species zhengi Gao, Y et Bu, Y, 2022:532

Multivesicula zhengi from China
After Gao Y & Bu Y, 2022 Fig.1



Genus Pongeiella Rusek, 1991

Species falca (Christiansen, K & Bellinger, P, 1980:490) Rusek, 1991, t.t.

  Subspecies europaea Rusek, 1991

             Syn.: Pongeiella falca ssp. europea(sic) Bu, Y et Gao, Y, 2019:71 ISS

Species luciaelvirae Thibaud & Peja, 1996

Species silvicola (Folsom, JW, 1932:57) Simón-Benito, JC, 2006:il


            Syn.: Tullbergia granulata Mills, HB, 1934:37
            Syn.: Tullbergia (Tullbergia) granulata Christiansen & Bellinger, 1992
            Syn.: Tullbergia granulata Christiansen, K et Bellinger, P, 1998:524
            Syn.: Paratullbergia granulata Simón-Benito, JC, 2006:il
            Syn.: Pongeiella granulata Simón-Benito, JC, 2006:il
Remark by Simón Benito, J.C.. 2006.10.16: This species should be revised, it does not belong in Mesaphorura (pseudocelli are not star shaped), but in Paratullbergia or Pongeiella.



            Syn.: Pongeiella silvicola Simón-Benito, JC, 2006:il
Remark by Janssens,F. 2024.09.13: Simón-Benito (2006) (personal communication) placed granulata Mills, 1934 in Pongeiella. But Christiansen & Bellinger (1992) already sunk granulata to silvicola Folsom, 1932. Therefore defacto Simón-Benito (2006) moved silvicola to Pongeiella.
Pongeiella silvicola from the USA
2024.08.26 © Wang, G.
Pongeiella silvicola juv. from the USA
2024.08.26 © Wang, G.
Pongeiella silvicola juv. from the USA
2024.08.26 © Wang, G.
Pongeiella silvicola juv. from the USA
Abd.6 granulation in transverse line
2024.08.26 © Wang, G.
Pongeiella silvicola juv. from the USA
Post-antennal organ with 2 rows of vesicles and outer part with 3 rows
2024.08.26 © Wang, G.



Species stojanovorum Pomorski & Skarzynski, 1997

Species yinchuanensis Bu, Y et Gao, Y, 2019:69


Genus Psammophorura Thibaud & Weiner, 1994

Species gedanica Thibaud & Weiner, 1994

             Syn.: Psammophorura gedanicus(sic) Bernard, EC, 2016:467 ISS

Species miniclavata Bernard, EC, 2016:464

Psammophorura miniclavata from the USA
After Bernard EC, 2016 Fig.8AB

Species neocaledonica Thibaud, J-M et Weiner, WM in Najt, J et Matile, L, 1997:77

Psammophorura neocaledonica from New Caledonia
After Thibaud J-M & Weiner WM, 1997 Fig.42


Genus Rotundiphorura Rusek, 1991

Species habanica Rusek, 1991


Genus Weinera Thibaud, 1993

Species ghislaineae Thibaud, 1993


Genus Sensilatullbergia Thibaud & Ndiaye, 2006

Species senegalensis Thibaud & Ndiaye, 2006


Genus Mexicaphorura Palacios-Vargas, JG et Catalán, E, 2013:92

Species guerrerensis Palacios-Vargas, JG et Catalán, E, 2013:93, t.t.

             Syn.: Mexicaphorura guerrensis(sic) Palacios-Vargas, JG et Catalán, E, 2013:94 ISS



Genus Ameritulla Bernard, EC, 2016:452

  Syn.: Collembola Amaritulla(sic) Miranda-Rangel, A, Cano-Santana, Z, Castaño-Meneses, G, Palacios-Vargas, JG et Corona-Tejeda, P, 2024:2607 ISS

Species clavata (Mills, HB, 1934:38) Bernard, EC, 2016:454, t.t.

Ameritulla clavata from the USA
After Bernard EC, 2016 Fig.1AB

Species hades (Christiansen & Bellinger, 1980) Bernard, EC, 2016:453

Species obscura Bernard, EC, 2016:458

Ameritulla obscura from the USA
After Bernard EC, 2016 Fig.4AB


Genus Mixturatulla Bernard, EC, 2016:461

Species ozwini Bernard, EC, 2016:461, t.t.

Mixturatulla ozwini from the USA
After Bernard EC, 2016 Fig.6ABC


Subfamilia Austraphorurinae Deharveng L, 2004:427

  Syn.: Tullbergiinae Austraphorurini Luciáñez & Simón, 1992



Genus Doutnacia Rusek, 1974

Species sp. Fjellberg A in Pomorski RJ & Skarzynski D, 1998:252

Species coineaui Massoud & Thibaud, 1985

Remark by Dunger, W. & Schlitt,B. 2011:35: This species probably belongs to another genus. Further characters separate the species from the genus Doutnacia. We consider the species a mere species inquirenda.



Species mols Fjellberg, A, 1998:146

            Syn.: Doutnacia ammophila Pomorski, RJ et Skarzynski, D, 1998:248


Species xerophila Rusek, 1974, t.t.

             Syn.: Doutnacia xerotherma(sic) Russell, DJ, 2008:124 lapsus ISS



Genus Jevania Rusek, 1978

Species fageticola Rusek, 1978, t.t.

Species weinerae Rusek, 1978

             Syn.: Jevania weineriae(sic) Busmachiu, G, 2010:153 ISS



Genus Scaphaphorura Petersen, 1965

Species arenaria (Petersen, 1965) Ellis, WN & Bellinger, PF, 1973:49, t.t.

Species corpesiensis Simón & Lucianez, 1990

Species cubana Thibaud, 1994

Species pakieti Pomorski, RJ et Skarzynski, D, 1997:3

Species sp. Dunger, 1977:181, s.i.


Genus Najtiaphorura Weiner & Thibaud, 1991

Species dobrolubovae Shveenkova, YB et Potapov, MB, 2003:?

Species marocana Thibaud & Boumezzough, 2006

Species olivieri (Weiner & Thibaud, 1991) Dunger, W & Schlitt, B, 2011:101

Species sandrinae Weiner & Thibaud, 1991


References